By Thomas S. Ray jr.

Slow-motionworld
of plant'behavior'
visiblein rain forest

Tiopical aines change their forms-

and the ir strat egi e s-s om etime s
euen reaching out for the darkness

Sheer diversity and a degree of adapta-

tion unimagined at temperate latitudes
help produce the stunning effect a tropi-
cal rain forest-the jungle we fantasize-
has on the first-time observer. Panama,
for example, is smaller than South Caro-
lina, yet contains more species of trees
(and birds and many other groups) than
all of North America north of Mexico.
The high diversity is qualitative as well
as quantitative. Adaptations occur in the
tropics that simply do not occur in tem-
perate zones. These adaptations, some-
times unexpected, sometimes bizane, in-
evitably provoke the curiosity of the re-
searcher. During several years of work
in the rain forests of Costa Rica, my own
curiosity was aroused by the adaptations
of vines-adaptations so unusual as to al-
ter my thinking about plants.
In the tropics vines are ubiquitous, and
have rarely been better described than by
Henry Walter Bates in 1863: "The tree
trunks were only seen partially here and Strands of monkey ladder vine mimic Costa Rica rain forest. Ruffiy bark of
there, nearly the whole frontage from sculpture as they reach toward light in vine supposedly makes it easier to climb.
ground to summit being covered with a
diversified drapery of creeping plants, all
of the most vivid shades of green; scarcely
a flower to be seen, except in some places how vines succeed in the most comPeti serted that plants are distinguished from
a solitary scarlet passion-flower, set in the tive environment on Earth. animals by not having the power of move-
green mantle like a star." It is my contention that, to better ex- ment. It should rather be said that plants
Since 1974 my colleagues and I have ploit their fellow plants, vines have acquire and display this power only when
tried to unravel the life histories of these evolved an elaborate behavioral reper- it is of some advantage to them; but that
specialized plants and their relations to toire. Few biologists consider Plants to this is of comparatively rare occurrence,
the rain forest around them. Starting exhibit any behavior at all, and with good as they are affixed to the ground, and food
with fragile seedlings in the litter of the reason. The notion of behavior is closely is brought to them by the wind and rain."
forest floor, we have followed them into linked to movement, something rare in A vine siarts its life as a seed, which
the canopy 80 to 100 feet overhead. Re- plants. However, it cannot be said that may be dispersed by a bird or bat, but
turning several times to Costa Rica, we plants are incapable of movement. As which germinates on the ground. It ma-
have become accustomed to trusting our Clrarles Darwin pointed out in On the tures and fruits, completing its life cycle,
lives to a rope, joining the birds and Mouements and Habits of Climbing in the top of a tree where light levels are
epiphytic orchids in the treetoPs to see Plants: "It has often been vaguely as- high. The tree is often far from the site

Sv..i f hson io.n

t2l
11a.r.!r l97g
of germination; the Plant may well have Dark Panel the black panel would be darker, seed-
spent most of its life wandering through lings would be attracted in that direc-
the forest in search of a suitable site for tion; but they would also seek to avoid
flowering. the bright light source itself, and so
Aroid vines (those in the familY of would grow off at an angle toward the
Araceae) are characteristically unbranch- dark side of the arena. Alternatively, the
ed; the stems are linear in shape. They seedlings could grow directly away from
grow at one end while gradually dying off the light source, or directly toward the
at the other as a result of senescence.In dark panel.
an accelerated time frame they would Therese Gurski, a junior at Lourdes
appear somewhat like green leafy snakes High School in Chicago, has completed
moving through the forest' But there is the experiment as a science fair project,
something very difierent about the move- under my direction. Her results show un-
ment of the plant "snake." While the ambiguously that the Monstera gigantea
body of the plant becomes displaced over seedlings headed straight for the dark
time, no part of the plant actually moves In this diagram of vine experiment, panel, almost as if they had a mechanism
(there is reason to believe, however, that seedlingshead straight for dark panel, for focusing the light, and could "see"
some nutrients are removed from senes- C. They did not simply grow away where they were going. To find a tree in
cent leaves and transported through the from brightest light, as in A, nor did the forest, the seedlings must Srow to-
stem to be used in the formation of new they avoid the average lightest area, as in ward discrete sourcesof darkness.
Ieaves at the growing end). B. Most plants grow strai8ht toward light.
Mobility in any organism creates the The philosopher class
need for orientation. Vines need an effi-
cient way of getting from their site of I have only been able to find one refer-
germination to the sunlight high in the to a meter in length as they search for a ence to the kind of movement that I have
trees. This can be thought of as a two-steP tree, The seedlings may occur in great observed,in a passageby Ambrose Bierce,
process.First the vine must encounter the densities around a tree bearing a mature a dialogue on whether machines-and
tree, and only then may it climb up into plant, and will all grow toward the tree, plants-can think. The protagonist asks:
the light. The former Processis the more appearing like short green spokes of a "'And what, pray, does it la machine]
interesting, as the latter can be effected wheel against the brown leaf litter, with think with-in the absence of a brain?'
by simply clinging to a tree and growing the tree like an axle joining the wheel at "The reply coming with less than his
up the trunk. the hub. customary delay, took his favorite form of
To understand the problem of finding Skototropism has a very short etymol- counter-interrogation :
a tree, imagine yourself to be a seedling ogy, beginiring in 1974 when Donald R' " 'With what does a plant think-in the
on the shady forest floor. Looking around, Strong jr. and I discovered the phenome- absenceof a brain?'
you see the light-green canoPy above, and non in Costa Rica. A tropism is an or- "'Ah, plants also belong to the phi
silhouetted against this are the long dark ganism's characteristic of growing in the losopher class! I should be pleased to
tree trunks. The key word here is dark. direction of something it needs, such as know some of their conclusions; you may
An efficient way of Setting to the nearest light or water. But this tropism was so omit the premises.'
tree is to grow toward darkness, a re- difierent from any other that we felt the "'Perhaps,' he replied, aPParently un-
sponse called skototropism that is very need to give it a new name. We finallY afiected by irony, 'you maY be
unusual for organisms that dePend settled on skotos, the Greek word for able to infer their convictions from their
directly on solar energy. Skototropism in "darkness, gloom" and formed the word acts. I will spare you the familiar ex-
plants is well developed only in vines, skototropism-growth toward darkness. amples of the.sensitive mimosa, the sev-
and it expressesitself only when the vine The phenomenon is not simPlY nega- eral insectivorous flowers and those whose
is "searching" for a tree' Once the tree is tive phototropism, growth away from stamens bend down and shake their
found, the plant switches back and starts light. This can be shown bY a simPle pollen upon the entering bee in order
growing toward light. experiment in which vine seedlings are that he may fertilize their distant mates.
Skototropism can produce a sPectacu- placed in the center of a circular arena But observethis. In an open spot in my
lar visual efiect on the forest floor' .labove). The arena consists of a wall of garden I planted a climbing vine. When
Monstera gigantea fruit contain thou- white cardboard. At one place on the wall it was barely above the surface I set a
sands of seeds,and when the fruit ripens, a black felt strip is attached. At an angle stake into the soil a yard away. The vine
the seeds are scattered on the ground of 90 degrees from this black strip, the at once made for it, but as it was about to
beneath the parent plant' The seedsger- wall is cut away and replaced by a trans- reach it after several days I removed it a
minate immediately, forming slender lucent panel; a light is placed behind it. few feet. The vine at once altered its
green leafless sprouts which can grow uP I can conceive of three difierent direc- course,making an acute angle, and again
tions the seedlings might grow. The made for the stake. This manoeuvre was
A graduate student at Harard, the plants might, in effect, take account of repeated several times, but finally, as if
author has been studying the ecolog the light coming from all directions. Be- discouraged,the vine abandoned the pur-
in Costa Rica or years. cause the side of the arena containing suit and ignoring further attemPts to
of rines f fiae

r22
 Build
Outdoor Sporting Specialtie s a museurn prece divert it traveled to a small tree. further
away, which it climbed.'
you'll be pnoud " 'And all this?'
" 'Can yow miss the significance of it?
to display. It shows the consciousness of plants. It
Send for our latest FREE CATALOG packed proves that they think.' "
with beautiful wooden ship model kits, can- The movement of the leafy "snake"
non kits, doll house furniture kits, model
books,tools, conslruction materialsand plans.
creates an interesting problem. The plant
grows acrossgreat distances, encountering
JAMESBL|SS& CO.,rNC.
widely varying microenvironments, from
MASS.02026
Dept.Sl, DEDHAM,
the cool dark of the forest floor where
debris falling from the reetops creates a
Malne Wlndfammer high risk of mortality, to those treetops
Sailing Vacation where the fresh air and bright tropical
CoastingSchoonas sunlight provide a benign atmosphere.
Isaac H. Evans The plant must be adapted to all of the
L,ewisR. French
$285.weekly. conditions that it encounters.At the same
BrochuresCAPTAINSLEE & FOSS,Box4827 time, every part of the plant remains
Rockland.Me. 04E41 Tel.207-594-8fi)7 fixed and thus must be highly fit in what-
ever surroundings it finds itself. A vine
Exotic Bitds Nstecards might adjust to this life history by evolv-
Finely lithoqraPhed notecards ing a form that is universally adapted,
iature colorful photos of ex.
otic waterbirds in natural habl- having a high fitness in the various micro-
Spring Catalog tats. Cards are folded, blank in-
side, brief descriptlon on back,
environments. Or it might evolve flexi
gift boxed, 12 cards-env./box, bility, so that it may express a difierent
Fullyillustrated. quality
Features apparel 2 ea. ol 6 birds. Two sets offer- form in each habitat. Thus each part of
andfootwear manand
fortheoutdoors ed. Set A or B, $6. Both sets,
the vine would assume the form which is
woman, leatherandcanvasbags,
furnishings $11. Shipplng, $1.0o. Pifron

forhome 0r camp, summer equip-

sports Pu bl ish ing, Box I 691-S, Sedona, best designed for the microenvironment
mentforfishing, camping,
canoeingand in which it must live. The latter aPProach
backpacking. All itemsguaranteed
to be is what we actually see in tropical vines.
10070 satisfactory.

For67years L. L. Beanhasprovided high

An AlternatlveXb
grademerchandise
personally
testtheproducts
prices.
at reasonable
weofferto
We Tlrc$21ButtonDown
thattheyrepresent
insure
practical,
functional
a truevalue,are
andhighin quality.
0urs...$14.
Weusefine,natural anddurable materials
suchasvirginwool, softcottonandcotton
fullgrained
flannels, leatherandsheepskin.

L.L. Bean is rugged

clothing enough to
withstand useyetattractively
outdoor styled
andcomfortable
forcasual wear.
0urfoot
weariscarefully
selectedto bewellfitting
andlongwearing.Manyitemsaremanu- ill'ift* coi,reeo
particularly COiTON OXFORD
factured
byusin Freeport, our BUTTONDOWN
shoes, slippers
boots. andbags.
Finesl traditional design and tailoring. Seven
button front, placket sleeve, lett breast pocket,
Youwillfindourservice personaland double reinforced side seams and sleeves.
Your satislactionguaranteed.
Foryourconvenience
efficient. ourtelephone Bl ue or W hi te. Si z es .14b17W 32- 35
order andretail
department inFreeport
store
areopen24hours a day,365daysa year. Illuntingtill @lolhirrs
2206BR YD ENR D .C O LU M BU S,OH IO43209
Pleasesend me 100o/o cotton shirt(s)@ $14.00ea
Add $1.75for s hj p. & Ins . r egar dl es sof quanti ty .
Ohio Res .add 47os al estax .
Size /- -White - Blue
Oly Qly.

tr Pers. Ck. ! Mstr. Chg. f] Visa n Am. ExP

Card No

Exp. Da te- N ,l s tr . C hg. Bank N o.- '-

Sig natu re
State- lip- Author's colleague, Don Strong, shoots
light line over a branch in tall tree.
L. L. Bean, Inc.
l0ll CascoSheet
Freeport, ME 04033
r-State

r24
 Are BirdsYour Thing?
As the result of an agreementrecentlyconcludedbetweenthe Smithsonian
Institution and the Yale University Pressto collaborateon the preplrration
and publicationof future volumesof Dr. Ralph W. Palmer'sauthoritative With climbing ropes in place, author
Handbook of North American Birds, the SmithsonianInstitution Pressis pulls himself up toward the canopy,
now able to offer to SmithsonianAssociatesat a specialprice the first
three publishedvolumesof this important series.
Regular Price AssociatesPrice To really understand how this works
Volume L, Loonsthrough Flamingos $32.sO $25.00 we must look closely at how vines are con-
structed. Aroid vines are composed of
Volume 2, Waterfowl (Part 1) $32.50 $2s.00
segments placed end to end. The seg-
Volume 3, Waterfowl (Part 2) $32.50 $25.00
ments consist of a piece of stem with a
$97.50 $7s.00 leaf and a pair of rootlets attached at the
forward end. The segmentsare produced
Savean additional$s.so by orderingall three volumesnow at $69.50. one at a time, at the grorving end of the
vine. The precise size and shape of each
Here are somenoteworthy reviews on Volume 1 segment can vary widely in response to
the type of environment in which it is
"A tightly edited, telegraphiccompilation of all that is significant about
being formed.
North Americanbirds, north of Mexico. . . ." Audubon Magazine
As a rule, vines make thicker segments
"The Handboolcseriesis destinedto becomea work-horseof professional in better environments. The ground is
ornithologiststhroughout the world and of all amateurbird watcherswho clearly the worst environment in the rain
do their'watching'in North America." Natural History forest for a plant, becauseof the low light
levels and falling debris. In Monstera
gigantea the terrestrial segments are the
The SmithsonianInstitution Press,P.O.Box7579, Washington, D.C. 20013. smallest. The lower part of tree trunks is
Pleasesend me the Handbook of North American Birds as indicated below: a habitat of intermediate quality, A vine
_Volumes 7,2, and 3 $69.50 on a tree has escapedmost of the danger
7 25'oo from falling debris, but on the lower part rtlNl
-volume
-Volume 2 25.OO of the trunk the leavesare still shaded by

fl
_Volume 3 25.00 the many layers of vegetation above. Here
-
S.rbtotul t}l.e Monstera assumes medium stature.
If the vine reaches the higher part of
Pleaseadd $7.50for postageand handlingfor the
the tree trunk, it has found the best con- rl
first book and 50 centsfor eachadditionalbook
ditions. In the presence of full sunlight ili .r
TOTAL-
and fresh air, and with the support of a
n Bill me ! Payment enclosed sturdy tree trxnk, Monstera gigantea will
undergo a transition in form that reflects
Name
its name. The stem grows and the leaves
Add
expand to a length o{ two meters, com-
City State - Zip -

r26
Monstera gi,ganteaseedlings, left, head for the tree
trunk in background. Climbing the tree, center,
pletely changing their shape and coming
to resemble enormous fronds (above)'
This behavior is understandable in that a
plant that is capable of moving from a
poor to a good environment would not be
likely to invest its precious resources in
constructing a full-sized plant body in the
worst of environments.
Variation in shape can be understood
even better by looking at the gnus Slrr-
gonium. The shape of the leaf is more
uniform, but the significance of changes
in the relative size of leaf and stem is
more obvious. Two segments of exactly
the same weight may look totally differ-
ent: one with a long stem and small
leaves, the other with a short stem and
very large leaves.
The first lorm, the long stem, is one
adapted almost entirely for "movement."
A segment with 95 percent of its material
in a long stem and only a vestige of a
leaf is barely able to photosynthesize,but
the plant has moved forward a distance
equal to the length of the stem. A segment
which has placed 95 percent of its ma-
r28
they develop small leaves. At the top of the tree,
vine produces frondlike leaves and fruit (right).
terial in a large leaf, with an extremely
short stem separating it from the previous
lea{, has created a large apparatus for
gathering light, while the plant has
moved forward only slightly.
In a good environment, Syngonium
segments naturally specialize for "feed-
ing"-and for immobility. They put most
of their sutrstance into large leaves not
only to take advantage of abundant sun-
light, but to reduce forward motion to a
minimum so they will not carry them-
selvesout of the good environment.
Segmentsin a poor environment have
a more interesting problem, They must
move, but they also have to "feed" them-
selves. So several contiguous segments
will take on the leafy {orm to gather what
sunlight there is. The next group assumes
the long-stemmed form, to make the
maximum forward progress. The Pattern
cycles back and forth until the vine en-
counters a tree.
This division of labor is repeated when
a vine climbs a small tree, only to find it
is shaded by taller trees. When the top of
the tree is reached, the plant has not yet
found the high light levels that it needs
to fruit. Generally the vine branches at
this point. One stem will remain in the
ffee while the other returns to the ground
in searchof another tree. The stem in the
tree continues to make relatively large
leafy segments, while the stem that has
returned to the ground makes long
slender leaflesssegments.The stem in the
tree is carrying on the photosynthesis
needed to support the stem searching for
another tree. If the searching stem is
broken ofi from the stem in the tree, it
will produce its own leaves, and behave
like the free living terrestrial form de-
scribed earlier.
This example illustrates how a mod-
erately large arboreal vine can be trans-
formed to a smaller terrestrial form
similar to the one it took much earlier in
its li{e. The development of vines is not
unidirectional. The vine is not geneti
cally programmed to produce a progres-
sion of forms leading it directly from the
seedling to the mature fruiting plant.
 IlreWndow
andGlidingDoor
AnswerBook
dndesed\d,lndwalle av
A poisonous snake,caught in a loop,
unexpectedly joined author in treetoPs.
Rather, the series of configurations ex- behavior.
pressed is entirely dependent on the se-
quence of environments that the vine
encounters as it moves through the rain
forest. No two individuals will show pre- the argument will persist until we
cisely the sameseriesof shapes.
as animals. There is a division of opinion
on the matter, and the line is not drawn
neatly between botanists and zoologists,
but rather along philosophical grounds.
There are those who feel that behavior
requires the existence of a nervous sys-
tem, an adaptation that is unique to the
animal kingdom.
On the other side there are those who
believe that plants have been misunder-
stood by us animals-that we lack sen-
sitivity to the ways of plants. For ex-
ample, awareness, in humans at least,
takes place on a much faster time scale
than the behavior described in this ar-
ticle. In an accelerated time frame one
could not help but notice a vine's highly
oriented growth movements, and the
changes in form in response to environ-
mental qualities. One would surely be
compelled to categorize such actions as
Ultimately, the semantic question can
be settled by agreement on a carefully
thought-out definition of the term. Yet
broaden our ways of thinking in an effort

Jo'PS'.*i:$l-i:$ "Genetic individuals" are capable of to understand the nature of plants. Per-

;sN;$Bis*Nlt:: being broken up into many independent

"physiological individuals," each of
which will have a different history. Thus
haps then we may even admit them to the
"philosopher class."

the fate of a seedling can be quite com-

plex. Even after reaching a large mature
fruiting form, an individual may become No longer an arboreal species,biologists
transformed into a small terrestrial plant use safety devicesin casethey fall.

*lffffili'-t'il1'--o""
resemtrling a seedling, This may happen
when the plant grows off the top of the
tree and returns to the ground as de-
scribed before. It may also happen if the

;l.$}try%;;'
vine is blown out of the treetops by a
storm or falls with a broken tree branch.
After being moved to the forest floor,
subsequently produced segmentswill take
on the configuration appropriate for that
environment. The vine retains complete
flexibility of form throughout its life.
In short, vines move through their rain

*tl*i*; forest habitat with a high degree of

orientation, "searching" for trees to climb
and then "seeking" light. As they en-
counter different environments, they
\\ttt'' assume difierent shapes. The changes in
form directly afiect the amount of photo-
D'&* synthesisthat takes place. Thus the plant
remains finely tuned to its environment
and maintains a high fitness throughout
;'4t.* its journeys.
Is it appropriate to speak of "plant be-

W"s ' *x )( on) D s hr AnJ er .en C .r n . gr !t.r t, V\ l 9'c

havior?" This interesting semantic ques-
tion impinges on how we see the world

Add;tional
rendong
ctry/srate/zip
q(peditions also !q: I Antarcljga ! Patagonia
LI Galapagos ls. Ll Mongolia Ll Easter lsland
FAMILY RINGS
GREST
1 4 K Go ld . $225'
Ste r lin g Silve r , t9 9 ;
WE WORK IN T HREE usGS(p.40)
BASIG WAYS- lf yo u Exploration and Empire' The Ex'
have a crest, send a
copy of it, otherwise plorer and the Scientist in the
yo u m a y ch e ck it Winning of the American West bY
a g a in st o u r list, o r yo u
m a y d e sig n Yo u r o wn William H. Goetzmann, KnoPf, 1966
(s e e b e l o w ) . S E N D F OR OUR F REE COL OR Goaernment in Science; The U.S.
CA T A L O G U E . l f . i n a d d itio n to fr e e ca ta lo g u e ,
y o u w a n t o u r b o o k l e t with o ve r 2 O,OOO names Geological Suraey, 1867-1894 bY
for which we have crests available plus in- Thomas G. Manning, UniversitY of
s tr u c t i o n s o n h o w t o d e sig n yo u r o wn cr e st,
sond us $1.OO. Kentucky Press (Lexington), 1967
HERATDICA INC.
IMPORTS, DOSM "A Brief History of the U.S. Geological
21 W E S T 4 6 t h S T . o NEW YORK, N.Y. 1 0 0 3 6
Suruey" by Mary C. Rabbitt, U.S.
Government Printing Office (Wash-
ington, D.C.), revised 1979
"A Guide to Obtaining Information
from the USGS 1978" by P E Clarke,
H. E. Hodgson and G. W. North,
USGS circular 777
Rodeos (p.56)
Man, Beast, Dust by Clifford P Wester-
meier, World Press (New York), 1947
Rodeo by Douglas Kent Hall, Ballan'
tine Books. 1976
Cow Country by Edward Everett Dale,
University of Oklahoma Press, 1942
Horses and Heroes by Frazier and
Robert Hunt, Charles Scribner's
Sons,1949
Kathakali (p. 68)
Dance Dialects of India by Ragini
Devi, Vikas (Bombay, London), 1972
Kathakali by Clifiord R. Jones and
Betty Tiue Jones, American Society
for Eastern Arts (San Francisco),
1970
The Dance in India by Enakshi Bhav-
nani, D. B. Taraporevala Sons & Co.
A full color Barbary Coast theatrical poster repro- Private Ltd. (Bombay), 1965
ducedfrom the archivesof the CaliforniaHistorical
So c i e t y b y t h e K o r be l F u n d . Yo u r s fo r $ 4 .9 5
(includespostageand handling).Posterrs 24" x35". Pheromones (p.78)
Make check payableto: Pheromones by Martin C. Birch, Am-
KORBEL FUND,BoxS
c/o CaliforniaHistoricalSociety
2090 JacksonStreet
SanFrancisco, California94109
r32
sterdam: North-Holland Publishing
Co.,1974
Race by John Baker, Oxford Univer-
sity Press(New York), 1974
"Human Olfactory Communication"
by Richard P Michael, Nature, April
8, 1976
"Menstrual Synchrony and Suppres'
sion" by Martha K. McClintock,
Nature,January22, l97l
Scarlet Letter (p. 86)
The ScarletLetter by Nathaniel Haw-
thorn, Hart Publishing Co', 1976
"Window on the Future: Planning for
'Public Television in the Telecom-
munications Era" bY Hartford N.
Gunn jr., Public Telecommunica'
tionsReaieu,July/August 1978
Black Confederates (p. 94)
Southern Negroes,1861-1865 by Bell
Irwin \44ley,YaleUniversity, 1988
A Diary from Dixie by Mary Boykin
Chesnut,D. Appleton, 1905
The Negro in the Ciail War by Ben-
jamin Quarles, Russell & Russell,
1953
The ConfederateStates of America,
1861-1865by E. Merton Coulter,
LouisianaStateUniversity,1950
Cities (p. 102)
Working Places:The Adaptiae Use ol
Industrial Buildings by Walter C.
Kidney, Oak Park Associates,Inc.
(Pittsburgh),1976
Inner City Historic Preseruationby
Arthur P Ziegler jr., The Allegheny
Press(Pittsburgh),l97l
Neighborhood Conseruation:Lessons
from Three Cities by Phyllis Myers
and Gordon Binder, The Conserva-
tion Foundation (Washington,D,C.),
r977
N eighborhoodConserttation : A Source
Booft,,editedby StephenA. Kliment,
The Whitney Library of Design
(New York), 1976
Vines (p. l2l)
The Power of Moaementin Plants bY
CharlesDarwin, D. Appleton, 1888
"Evolutionary Significance of Pheno-
typic Plasticityin Plants" by A. D.
Bradshaw, Aduances in Geneti'cs,
(Vol. l3), 1965
"Seedlingsin Searchof the Dark" by
Donald R. Strong ir., ScienceNews,
November29, 1975